THE CONSERVATION OF THE - ULSpace

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DECLARATION

I declare that the dissertation hereby submitted to the University of Limpopo for the degree of Master of Science has not previously been submitted by me for a degree at this or any other university, that it is my own work in design and in execution, and that all material contained therein has been duly acknowledged.

_________________________

Joseph I. Grosel (Candidate)

Date: ____________________

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PREFACE

The study was carried out through the Department of Biodiversity in the School of Molecular and Life Sciences, University of Limpopo from January 2005 to October 2007, under the supervision of Professors S.M. Dippenaar and G.D. Engelbrecht. This study represents original work by the author and where the work of other authors has been used they are duly acknowledged in the text and listed as references.

Chapter 1 consists of a general introduction in which the broad characteristics and taxonomy of the Alaudidae family are discussed. The southern African representatives of the family are introduced with particular emphasis on the Short-clawed Lark Certhilauda chuana and the objectives of the study are outlined. Chapters 2 to 4 of this dissertation have been written in the form of research papers with the relevant tables and figures appearing at the end of each chapter. In chapter 2 the taxonomic status of the eastern population of the Short-clawed Lark is clarified by comparing molecular, vocalisation and morphometric data with that of the western population. Chapter 3 describes the distribution, population status and habitat preference of the eastern population of the Short-clawed Lark while chapter 4 details the vocalisations and territorial display of the species. Chapter 5 is a culmination of the findings of the previous chapters and includes an assessment of its conservation status, threats faced by the eastern population, research and conservation recommendations.

Large tables appear as appendices and a single list of references for all five chapters is

included in the back of the dissertation. To give this manuscript a degree of uniformity,

the literature cited and associated citations in all the chapters have been formatted

according to the manuscript requirements of the Journal of African Zoology. Due to the

format of this dissertation, a certain amount of duplication of information has resulted,

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Relevant scientific publications, papers read and posters presented at congresses and symposia are listed below (* presenting author):

ENGELBRECHT, G.D.*, GROSEL, J.I. & VAN TONDER, R. Comparative growth curve analysis of two sympatric lark species: the Short-clawed and Sabota Lark. Poster presented at the IV^th Natural Forests and Woodland Symposium, Port Elizabeth, South Africa - May 2006

GROSEL, J.I., DIPPENAAR, S.M.* & ENGELBRECHT, G.D. The Lark and the Short of it: The status of the Short-clawed Lark in South Africa. Poster presented at the 5^th South African Society for Systematic Biology Congress, Kruger National Park, South Africa – July 2006.

ENGELBRECHT, G.D.*, GROSEL, J.I. & DIPPENAAR, S.M. Concerning the “least concern”:

Conservation of the Short-clawed Lark in South Africa. Paper read at the 24^th International Ornithological Congress, Hamburg, Germany – August 2006.

GROSEL, J.I.*, ENGELBRECHT, G.D. & DIPPENAAR, S.M. The Biology and Ecology of the Short-clawed Lark (Certhilauda chuana). Paper read at the Environmental Research in Limpopo Congress, Polokwane, South Africa – November 2006.

ENGELBRECHT, G.D.*, GROSEL, J.I. & VAN TONDER, R. Conservation of the biodiversity of the Polokwane Plateau. Paper read at the 41^st Annual Congress of the Grassland Society of Southern Africa, Bela-Bela, South Africa. July 2006.

ENGELBRECHT, G.D., GROSEL, J.I. & DIPPENAAR, S.M. 2007. The Western Population of the Short-clawed Lark Revisited. Bulletin of the African Bird Club 14(1): 58-60.

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ABSTRACT

The southern African endemic Short-clawed Lark Certhilauda chuana comprises two geographically isolated populations, consisting of a western and an eastern population.

Several authors have suggested that the taxonomic status of the eastern population be verified. In an attempt to resolve the taxonomic uncertainty regarding the two populations, DNA molecular data was used in a comparative study of the two populations. The DNA sequence data was generated using standard methods of DNA extraction, PCR and automated DNA sequencing for partial sequences of the mitochondrial genes Cytochrome b and ND2. A total of 530 base pairs of the Cytochrome b gene were amplified for individuals representing both Short-clawed Lark populations. The results obtained from the amplified Cyt b sequences showed the populations to be identical. The amplified partial ND2 gene (972 base pairs) showed sequence divergence between the eastern and western populations ranging between 0.10 - 0.31%. The partial ND2 gene produced only four haplotypes for both the eastern and western individuals with a single mutational step separating each of the haplotypes.

The results of the DNA analysis showed that there exists very little genetic diversity within and between the two populations.

Confirmation of the taxonomic status of the two populations was supplemented by comparing selected morphometric measurements and territorial song characterization.

Apart from a statistically significant difference (P < 0.05) in the length of the tarsus, there

were no other significant differences in the morphometric parameters analysed. A

comparison of the territorial calls of males showed considerable individual variation

between males within a population and statistically significant differences between males

from the two populations for some of the parameters analysed.

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The results of a habitat preference study for the eastern population was determined by analysis of the micro- and macro-habitat features within presently and historically occupied territories. It was established that as with the western population, Short-clawed Larks from the eastern population also show a fine-scale habitat preference within the broad, general habitat description of “open habitat, sparsely vegetated with small trees and shrubs”. The results of the present study explain the highly localized distribution of the species within its area of occurrence. The results also suggest that the species’

habitat preference is probably dictated as much by physiological requirements, e.g.

short-grassed areas with bare ground for breeding and foraging, as it is by behavioural requirements, e.g. large open areas for aerial displays and small trees or shrubs for territorial calls.

In addition to the above, surveys were conducted to determine the species’ present extent of occurrence, area of occupancy and to obtain population size estimates. The results were compared with published data from the Southern African Bird Atlas Project and showed a dramatic range reduction of this species. Possible reasons for the range reduction include habitat loss, absence of formally protected habitat and altered ecological processes such as lack of fire and bush encroachment. It is estimated that the eastern population of the species comprise fewer than 380 breeding pairs. The observed range reduction and estimated population size suggests that the eastern population of the species requires urgent protection.

A comprehensive investigation of Short-clawed Lark vocalisations and displays was

conducted. This study presents the first detailed analysis of the vocalisations of the

Short-clawed Lark. Analysis of call data showed considerable variation in the number

and placement of pulsed and whistle notes in their different calls. The extent of individual

variation within the eastern population for the parameters analyzed, failed to reveal any

dialects associated with the vocalisations of Short-clawed Larks from the eastern

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population. Although Short-clawed Larks call and display throughout the year and during most parts of the day, most vocalisations and displays peak in the first 1-3 hours after sunrise in the peak-breeding season. This information is useful for identification and monitoring purposes.

This manuscript concludes with a discussion of the major threats facing the population,

recommendations on how to address them, and conservation priorities for the eastern

population and the species.

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ACKNOWLEDGEMENTS

I wish to acknowledge and thank the following people who assisted and advised me at various stages of this project, and the institutions for making this study possible.

Firstly, my two supervisors:

 Professor Derek Engelbrecht for his expert guidance, encouragement and

friendship throughout the study and for miraculously managing to convince a forty-year old to return to the academic field.

 Professor Susan Dippenaar, for her patience and proficient supervision in the lab

and for her willingness to offer experienced advice whenever it was required.

 The Department of Biodiversity in the School of Molecular and Life Sciences, of

the University of Limpopo who provided much needed funding in order to undertake fieldtrips to the North West Province and for laboratory expenses during the genetic analysis phase of the project.

 The Polokwane Municipality who allowed me unrestricted access into the

Polokwane Nature Reserve and various other municipal properties.

 The Pretoria laboratories of Inqaba Biotechnical Industries for their valuable

contribution with the genetic sequencing.

 The staff and curators of the National Flagship Institute in Pretoria and the

Durban and McGregor museums who provided museum study skins.

 Mrs. Rita Olwagen for offering her very kind assistance and expertise with the

statistical analysis of the study.

 Dr Ute Kryger from the Department of Zoology and Entomology of the University

of Pretoria for her proficient assistance with the analysis and interpretation of the

molecular data.

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 Mr. Rowan van Tonder, for his assistance in the field and for the tedious task of

transferring field data onto computer spreadsheets.

 Pete Laver who shared his time and GIS proficiency which resulted in the

production of the maps in this dissertation.



“Oom” Kalie Lotz and Mr. Maxwell Ledwaba from the Polokwane Municipality’s Environmental Management Business Unit who recognized the importance of the study and supported it by providing the relevant resources and encouragement.

 Mr. Ken Lund and Mr. Jan van Waveren who made their farms available for the

study.

I also wish to thank the following special people in my life.

 Uncle Kit and Aunty Katica Sutton whose affection and generosity made this and

previous tertiary studies possible.

 To my Mother for her unfailing love and to my late farther who, instilled in me, a

passion for all that is nature - this work is dedicated to him.

 Lastly, to my girlfriend, business partner and fiancé who all became my wife

during the course of this study, for all the sacrifices and her valuable support and

encouragement, particularly during the seemingly endless word processing phase

of the project.

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TABLE OF CONTENTS

Page

DECLARATION ……….. i

PREFACE ……… ii

ABSTRACT ………. iv

ACKOWLEDGEMENTS ……… vii

CHAPTER 1 - General Introduction ……….. 1

Larks (Family Alaudidae) ………..……….. 1

Characteristics of the family Alaudidae ………. 2

Alaudidae taxonomy ……… 5

Southern African members of Alaudidae ……… 11

The Short-clawed Lark, Certhilauda chuana ………... 12

Description ……… 13

Habitat, movement, and general habits ………... 14

Call and display ………... 15

Breeding ……… 15

Status, distribution and range ……… 16

Current study ………. 18

Objectives of the study ……… 19

CHAPTER 2 - The taxonomic status of the eastern population of the Short- clawed Lark Certhilauda chuana ……….. 21

Introduction ……… 21

Materials and Methods ……… 23

Morphometric Study ……… 23

Sampling ……….. 23

Morphometrics ……… 24

Territorial song characterisation ……… 24

Statistical analysis of vocalisations ………. 25

Genetic Study ……….. 25

Sampling of genetic material ……… 25

DNA extraction and Polymerase Chain Reaction (PCR)... 26

Cyt b ………. 26

ND2 ……….. 27

Sequencing ………. 28

Phylogenetic Analysis ……… 28

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Results ………... 30

Morphometric results ……….. 30

Territorial call analysis……… ……… 30

Molecular genetic analyses ………... 32

Sequence statistics ……… 32

Sequence divergence ……… 33

Phylogenetic analyses ……….. 34

Population genetic variability and population subdivision … 34 Phylogeographic analysis ………. 35

Discussion ………. 35

CHAPTER 3 - Distribution, population status and habitat preference of the eastern population of the Short-clawed Lark

Certhilauda chuana

in the Limpopo Province, South Africa ……….. 52

Material and Methods ……….. 54

The study area ………. 54

Ecological survey ……… 55

Site selection for ecological surveys ………... 55

Vegetation and habitat surveys ……… 56

Micro-habitat survey ……….. 57

Macro-habitat survey ………. 59

Statistical Analysis ………. 60

Survey of the extent of occurrence and area of occupancy 61 Results ………... 62

Ecological survey results ……… 62

Extent of Occurrence and Area of Occupancy……… 65

Discussion ………. 67

CHAPTER 4 - Vocalisations and aerial displays of the Short-clawed Lark

Certhilauda chuana ………..

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Materials and methods ……… 98

Vocalisations ……… 98

Frequency of singing and aerial display flights ……….. 99

Results ………... 101

Vocalisations ……… 101

Territorial calls ………. 101

Agitated calls ……… 102

Contact calls ………. 102

Nestling distress calls ………. 103

Frequency of singing and display ………. 103

Discussion ………. 104

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Display Flight ……… 107

CHAPTER 5 - Aspects of the conservation of the Short-clawed Lark

Certhilauda chuana, with special reference to the eastern population………..

118 Introduction ……… 118

5.1. Overview of current knowledge regarding the Short-clawed Lark …… 119

5.1.1. General biology of the Short-clawed Lark ……… 119

5.1.2. Habitat preference of the Short-clawed Lark ………... 120

5.1.3. Ethology with relevance to identification and monitoring ….. 120

5.1.4. Population differentiation and taxonomy ……….. 121

5.1.5. Extent of occurrence and estimated population numbers …. 122 5.1.6. Conservation status of the population and species ………… 123

5.2. Threats ………... 124

5.2.1. Habitat loss or change ………. 124

5.2.1.1. Recommendations ………... 127

5.2.2. Altered ecological processes ………. 127

5.2.3. Lack of formally protected habitat ………. 131

5.2.4. Direct and indirect anthropogenic effects ………. 134

5.2.4.1. Recommendations ………... 135

5.2.5. Predation ………... 136

5.2.6. Pesticides and herbicides ...……… 137

5.2.7. Stochastic and small population threats ………... 138

5.2.8. Climate Change ……… 139

5.3. Opportunities ………. 140

5.3.1. The role of “avi-tourism” ……….. 140

5.3.2. The role of NGO’S ……… 141

5.4. Research needs ……… 142

5.5. Proposed conservation priorities ……… 142

Conclusion ………. 143

REFERENCES ……… 149

APPENDIX……….... 163

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LIST OF TABLES, FIGURES AND APPENDICES.

TABLES.

Title/caption Page

Table 2.1. Details of Short-clawed Lark specimens used for morphometric measurements from the western (W) and eastern (E) populations.

Polokwane Nature Reserve = PNR (eastern population) and Botsalano Nature Reserve = BNR (western population).

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Table 2.2. Sampling localities for genetic analyses. Short-clawed Larks Certhilauda chuana from the eastern population (n = 6) (1E – 6E) and from the western population (n = 6) (1W – 6W). Outgroup birds consisted of Rufous-naped Lark Mirafra africana (n = 2) (1R & 2R) and Spike-heeled Lark Chersomanes albofasciata (n = 3) (1S – 3S).

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Table 2.3. Sequences and locations of primers used for amplification and sequencing of Cytochrome b and ND2 genes.

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Table 2.4. Results of morphometric comparisons between male Short-clawed Larks from the eastern and western populations (Mann-Whitney U test).

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Table 2.5. The number of single-, double- and triple-note territorial call phrases and their relative contributions recorded for Short-clawed Lark males from the eastern and western population.

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Table 2.6. Single and double-note territorial call variables analysed for Short-clawed Lark males from the eastern and western population of the species.

*Statistically significant (P < 0.05) differences between populations. a) Single-note territorial call variables - DT = delta time (ms), LF1 = Lowest Frequency (kHz), HF1 = Highest Frequency (kHz), DF1 = Delta frequency (kHz). (Variables with significant differences = P < 0.05) b) Double-note call variables - DT1, DT2 and DTTot = delta time of the first and second note and total phrase respectively (ms), LF1, LF2 and LFTot = Lowest Frequency of the first and second note and total phrase respectively (kHz), HF1, HF2 and HFTot = Highest Frequency of the first and second note and total phrase respectively (kHz), DF1, DF2 and DFTot = Delta frequency of the first and second note and total phrase respectively (kHz).

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Table 2.7. Pairwise estimates of ND2 nucleotide sequence divergence between Short- clawed Larks from the Western population (1W – 6W) and the Eastern population (1E – 6E) with outgroups, Rufous-naped Lark (1R & 2R) and Spike-heeled Lark (1S – 3S). Sequence differences are depicted in percentage.

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Table 3.1. Summary of mean grass height (height classes – 1. = 0 to 50mm, 2. => 50 to 150mm, 3. = >150 to 250mm, 4. = >250 to 500mm, and 5. = >500mm) and percentage basal cover in currently occupied (n = 40) and unoccupied (n = 20) territories of the eastern population of the Short-clawed Lark. (P = 0.05)

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Table 3.2. Grass species prevalence, measured in territories of the eastern population of the Short-clawed Lark (n = 40). The species richness was 57 species.

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Table 3.3. Woody plant species prevalence in territories of the eastern population of the Short-clawed Lark

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population of the Short-clawed Lark.

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Table 4.1. Sampling localities for analysis of vocalisations of territorial male Short- clawed Larks Certhilauda chuana from the eastern population of the species (n = 13).

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Table 4.2. The number of single-, double- and triple-note territorial calls recorded for Short-clawed Lark males from the eastern population of the species. The relative percentage contribution of each note to the total number of notes is presented in parenthesis.

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Table 4.3. Single-note territorial call (n = 49) variables analysed for Short-clawed Lark males from the eastern population of the species. DT = delta time (ms), LF1

= Lowest Frequency (kHz), HF1 = Highest Frequency (kHz), DF1 = Delta frequency (kHz).

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Table 4.4. Double-note territorial call variables analysed for Short-clawed Lark males from the eastern population of the species. DT1, DT2 and DT-Tot = delta time of the first and second note and total phrase respectively (ms), LF1, LF2 and LFTot = Lowest Frequency of the first and second note and total phrase respectively (kHz), HF1, HF2 and HFTot = Highest Frequency of the first and second note and total phrase respectively (kHz), DF1, DF2 and DF_Tot = Delta frequency of the first and second note and total phrase respectively (kHz).

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Table 4.5 Triple-note territorial call variables analysed for Short-clawed Lark males (n

= 32) from the eastern population of the species. DT1, DT2, DT3 and DT-Tot

= delta time of the first, second and third note and total phrase respectively (ms), LF1, LF2, LF3. LFTot = Lowest Frequency of the first, second and third note and total phrase respectively (kHz), HF1, HF2, HF3. HFTot = Highest Frequency of the first and second and third note and total phrase respectively (kHz), DF1, DF2, DF3. DFTot = Delta frequency of the first and second and third note and total phrase respectively (kHz).

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Table 4.6. Contact call variables (n = 171) analysed for Short-clawed Larks from the eastern population of the species. DT = Delta Time (ms), LF1 = Lowest Frequency (kHz), HF1 = Highest Frequency (kHz), DF1 = Delta Frequency (kHz), NN = Number of notes.

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Table 4.7. Nestling distress (n = 25) call variables of the Short-clawed Lark. DT = Delta Time (ms), LF1 = Lowest Frequency (kHz), HF1 = Highest Frequency (kHz), DF1 = Delta Frequency (kHz).

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Table 4.8. Sunrise and sunset times in the study area when call and display data was collected.

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Table 5.1. Summary of the biological characteristics of the Short-clawed Lark. 144 Table 5.2. A summary of the features that constitute the preferred habitat of the eastern

population of the Short-clawed Lark.

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Table 5.3. Evaluation of the status of the Short-clawed Lark based on IUCN Red List categories and criteria (2005). Values for the eastern population are based on the results of the present study while those for the western population are derived from published data.

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Table 5.4. Farms selected as important Short-clawed Lark areas where, as a result of certain land management practices, viable breeding habitat have been created for the species.

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Table 5.5. Conservation priorities for the eastern population of the Short-clawed Lark. 148

FIGURES.

Figure 1.1. Male (left) and female (right) Short-clawed Larks showing distinct facial features i.e. prominent cream-coloured supercilium, dark brown eye-stripe, light russet cheeks and ear coverts, streaked crown, dark eye and light base to lower mandible. Plumage staining as a result of dusting can also be seen on the upper flanks of the female bird.

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Figure 1.2. The coat-of-arms of the Polokwane Municipality featuring the Short-clawed Lark.

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Figure 2.1. A strict consensus tree of the most parsimonius trees of the partial Cytochrome b gene from individuals from the western (1W – 6W) and eastern (1E – 6E) populations of the Short-clawed Lark with outgroup individuals represented by the Rufous-naped Lark (1R & 2R) and the Spike-heeled Lark (1S – 3S).

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Figure 2.2. Most parsimonious tree with bootstrap values for the ND2 dataset (CI = 0.995, RI = 0.997) from individuals from the western (1W – 6W) and eastern (1E – 6E) populations of the Short-clawed Lark with outgroup individuals represented by the Rufous-naped Lark (1R & 2R) and the Spike-heeled Lark (1S – 3S). Numbers on the branches refer to bootstrap support values.

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Figure 2.3. The ND2 Midpoint-rooted bootstrap tree representing the ingroup individuals from the western (1W – 6W) and eastern (1E – 6E) populations of the Short-clawed Lark.

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Figure 2.4. TCS network of four haplotypes of individuals from the Eastern and Western Short-clawed Lark populations where C = the 4E singleton, D = the 5W singleton, A = the rest of the Eastern population individuals and B

= the rest of the Western population individuals. A also represents the ancestral population.

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Figure 3.1. Average monthly precipitation (1996-2006) for the greater Polokwane area recorded at the Polokwane/Gateway International Airport weather station - 0677802 BX (South African Weather Service, 2006).

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Figure 3.2. Schematic representation of site selection and survey methodology for macro- and micro-habitat analysis.

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Figure 3.3. The percentage of occupied territories (n = 40) and unoccupied sites (n = 20) of the Short-clawed Lark showing different degrees of slope.

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Figure 3.4. A representative sample of altitudes recorded in the territories of Short- clawed Larks from the eastern population.

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Figure 3.5. The percentage of Short-clawed Lark territories (n = 40) showing their 85

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Figure 3.6. Mean percentage ground cover in occupied territories and unoccupied sites of the Short-clawed Lark, where RO = Occupied lark territories within the (n = 20), OO = Occupied territories outside protected areas (n = 20), RU = Unoccupied sites in the Polokwane Nature Reserve (n = 10) and Unoccupied sites outside protected areas (n = 10).

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Figure 3.7. Mean grass height in occupied territories and unoccupied sites of the Short-clawed Lark. Grass height classes: Class 1 = 0 to 50mm; Class 2 =

>50 to 150mm; Class 3 = >150 to 250mm; Class 4 = >250 to 500mm and Class 5 = >500mm. The four site categories are; RO = Occupied lark territories within the Polokwane Nature Reserve (n = 20), OO = Occupied territories Outside protected areas (n = 20), RU = Unoccupied sites in the Reserve (n = 10) and OU = Unoccupied sites Outside protected areas (n = 10).

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Figure 3.8. Grass species dominance in a) occupied territories (n = 40) and in b) unoccupied territories (n = 20) of the Short-clawed Lark from the eastern population.

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Figure 3.9. Mean percentage tree cover in occupied territories (n = 40) and

unoccupied sites (n = 20) of the eastern population of the Short-clawed Lark.

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Figure 3.10. Mean percentage tree height and tree crown diameter within occupied territories (n = 40) and unoccupied sites (n = 20) of the eastern population of the Short-clawed Lark.

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Figure 3.11. The percentage of occupied territories (n = 40) and unoccupied sites (n = 20) of the Short-clawed Lark showing different degrees of grazing.

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Figure 3.12. The percentage of occupied territories (n = 40) and unoccupied sites (n = 20) of the Short-clawed Lark showing different levels of habitat succession based on grass species abundance.

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Figure 3.13. The percentage of occupied territories (n = 40) and unoccupied sites (n = 20) of the Short-clawed Lark showing the mean ecological status based on grass species prevalence and composition.

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Figure 3.14. The percentage of occupied territories (n = 40) and unoccupied sites (n = 20) of the Short-clawed Lark showing its fire history. Classes represent the history of fire occurrence prior to the survey date, categorized into 12- month time frames.

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Figure 3.15. The present area of occupancy of the eastern population of the Short- clawed Lark within the Polokwane Plateau Bushveld, Mamabolo Bushveld and Makhado Sweet Bushveld vegetation units (Mucina & Rutherford 2006).

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Figure 3.16 Locations of 57 occupied Short-clawed Lark territories recorded during the 2005/2006 breeding season within relevant vegetation communities of the Polokwane Nature Reserve.

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Figure 3.17. The distribution of the eastern population of the Short-clawed Lark during the South African Bird Atlas Project 1 (left) compared to the results of the current distribution survey (right).

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Figure 3.18 Population estimates of the eastern population of the Short-clawed Lark across its present area of occurrence.

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Figure 3.19 Photographs of suitable Short-clawed Lark habitat A) within the Polokwane Nature Reserve and B) and on the farm Snymansdrift 738LS, west of Polokwane City. Photograph C) shows Acacia hebeclada and A.

tortilis shrub encroachment in the rural areas of Mothibaskraal (east of Polokwane) as a result of deforestation and overgrazing.

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Figure 3.20 Photographs showing some of the phenotypic similarities in plumage and shape between the Short-clawed Lark Certhilauda chuana and the Sabota Lark Calendulauda sabota. Photograph A) - Short-clawed Lark above, Sabota Lark below. Photograph B) - Sabota Lark left, Short-clawed Lark right. Photograph C shows the distinct size sexual dimorphism of the Short-clawed Lark with the much larger male below.

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Figure 4.1. Sonogram of a triple-note Short-clawed Lark territorial call. 115 Figure 4.2. Sonogram of a typical agitated call by a male Short-clawed Lark. 115 Figure 4.3. Sonogram of a 3-note Short-clawed Lark contact call. 115 Figure 4.4a The mean frequency of territorial calls by male Short-clawed Larks over a

year.

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Figure 4.4b The mean frequency of Short-clawed Lark contact calls over a year. 116 Figure 4.4c The mean frequency of agitated Short-clawed Lark calls over a year. 117 Figure 4.5. Mean number of single display flights by male Short-clawed Larks over

four seasons.

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APPENDIX.

APPENDIX 1.1 A comprehensive list of the larks (Alaudidae) of the world based on taxonomy according to Sibley & Monroe (1993), Sinclair & Ryan (2003) and Hockey et al. (2005).

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APPENDIX 2.1 Nucleotide sequences of 530 base pairs of the mitochondrial Cytochrome b gene of Short-clawed Larks from the eastern (1E-6E) and western population (1W-6W) aligned with those of outgroups; Rufous- naped Lark Mirafra africana (1R & 2R) and Spike-heeled Lark Chersomanes albofasciata 1S-3S. Dots indicate identity to the 1E sequence. These sequences correspond to locations on the chicken (Gallus) sequence (Desjarins & Morais 1990) of the 3' end of the primer sequence.

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APPENDIX 2.2 Nucleotide sequences of 972 base pairs of the mitochondrial ND2 gene of Short-clawed Larks from the eastern population (1E-6E) and western population (1W-6W) aligned with that of outgroups; Rufous-naped Lark Mirafra africana (1R & 2R) and Spike-heeled Lark Chersomanes albofasciata 1S-3S. Dots indicate identity to the 1E sequence.

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APPENDIX 3.1 Field data sheets used in the collection of data for habitat preference study of the eastern population of the Short-clawed Lark.

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THE

SHORT-CLAWED LARK

Certhilauda chuana